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Physiology of Elasmobranch Fishes Internal Processes 1st
Edition Robert E. Shadwick Digital Instant Download
Author(s): Robert E. Shadwick, Anthony P. Farrell, Colin J. Brauner
ISBN(s): 9784584594728, 4584594724
Edition: 1
File Details: PDF, 44.25 MB
Year: 2015
Language: english
PHYSIOLOGY OF
ELASMOBRANCH FISHES:
INTERNAL PROCESSES
This is Volume 34B in the

FISH PHYSIOLOGY series


Edited by Anthony P. Farrell and Colin J. Brauner
Honorary Editors: William S. Hoar and David J. Randall

A complete list of books in this series appears at the end of the volume
PHYSIOLOGY OF
ELASMOBRANCH FISHES:
INTERNAL PROCESSES

Edited by

ROBERT E. SHADWICK
Department of Zoology
The University of British Columbia
Vancouver, British Columbia
Canada

ANTHONY P. FARRELL
Department of Zoology, and Faculty of Land and Food Systems
The University of British Columbia
Vancouver, British Columbia
Canada

COLIN J. BRAUNER
Department of Zoology
The University of British Columbia
Vancouver, British Columbia
Canada

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Designer: Matthew Limbert
CONTENTS

CONTENTS OF PHYSIOLOGY OF ELASMOBRANCH FISHES:


STRUCTURE AND INTERACTION WITH ENVIRONMENT, VOLUME 34A ix
CONTRIBUTORS xiii
PREFACE xv
LIST OF ABBREVIATIONS xix

1. Elasmobranch Cardiovascular System


Richard W. Brill and N. Chin Lai
1. Introduction 2
2. Cardiovascular Function and Energetics 3
3. Factors Controlling and Effecting Cardiovascular
Function 24
4. Signaling Mechanisms Effecting Blood Vessel Diameter 37
5. The Action Potential and Excitation–Contraction (EC) Coupling in
Elasmobranch Hearts: The Influences of Environmental, Biochemical,
and Molecular Factors 48
6. Practical Applications: Physiology in the Service of Elasmobranch
Conservation 54
7. Summary 57

2. Control of Breathing in Elasmobranchs


William K. Milsom and Edwin (Ted) W. Taylor
1. Introduction 84
2. Ventilation: Efferent Motor Output to the Respiratory
Muscles 85
3. Central Respiratory Rhythm Generation: The Source of the Motor
Output 91
4. The Respiratory Pattern: The Conditional Nature
of the Output 94

v
vi CONTENTS

5. Relationships Between Ventilation and Heart Rate 103


6. Afferent Input 108
7. Conclusions 115

3. Oxygen and Carbon Dioxide Transport in Elasmobranchs


Phillip R. Morrison, Kathleen M. Gilmour
and Colin J. Brauner
1. Introduction 128
2. Blood-Oxygen Transport 132
3. Transport and Elimination of Carbon Dioxide 184
4. Conclusions and Perspectives 197

4. Organic Osmolytes in Elasmobranchs


Paul H. Yancey
1. Introduction 222
2. Osmoconformers Versus Osmoregulators 224
3. Properties of Organic Osmolytes 233
4. Metabolism and Regulation 256
5. Evolutionary Considerations 263
6. Knowledge Gaps and Future Directions 264

5. Regulation of Ions, Acid–Base, and Nitrogenous Wastes


in Elasmobranchs
Patricia A. Wright and Chris M. Wood
1. Introduction 280
2. Ionoregulation 280
3. Acid–Base Balance 298
4. Nitrogenous Wastes 310
5. Concluding Remarks 327

6. Feeding and Digestion in Elasmobranchs: Tying


Diet and Physiology Together
Carol Bucking
1. Introduction 348
2. Feeding Habits of Elasmobranchs 350
3. Elasmobranch Gastrointestinal Tract Anatomy 365
4. Digestive Enzymes and Secretions 373
5. Effects of Digestion on Homeostasis 379
6. Future Perspectives 381

7. Metabolism of Elasmobranchs (Jaws II)


J.S. Ballantyne
1. Introduction 396
2. Evolutionary Context 398
CONTENTS vii

3. Diet and Digestion 399


4. Oxidative Metabolism 404
5. Carbohydrate Metabolism 407
6. Nitrogen Metabolism 411
7. Lipid and Ketone Body Metabolism 426
8. Vitamin Metabolism 435
9. Xenobiotic Metabolism 436
10. Conclusions and Perspectives 437

8. Endocrine Systems in Elasmobranchs


W. Gary Anderson
1. Introduction 458
2. Pituitary Gland 459
3. Corticosteroids and Catecholamines 472
4. Gastro-Entero-Pancreatic Hormones 484
5. The Heart as an Endocrine Gland 497
6. The Kidney as an Endocrine Gland 502
7. The Pineal 505
8. Calcium Regulation 506
9. Conclusions and Perspectives 508

INDEX 531
OTHER VOLUMES IN THE FISH PHYSIOLOGY SERIES 549
CONTENTS OF
PHYSIOLOGY OF ELASMOBRANCH FISHES:
STRUCTURE AND INTERACTION WITH
ENVIRONMENT, VOLUME 34A

1. Elasmobranchs and Their Extinct Relatives: Diversity,


Relationships, and Adaptations Through Time
Philippe Janvier and Alan Pradel
1. Introduction 2
2. Systematic and Phylogenetic Framework of Chondrichthyan Diversity 4
3. Environments and Adaptations 13
4. Conclusion 14

2. How Elasmobranchs Sense Their Environment


Shaun P. Collin, Ryan M. Kempster and Kara E. Yopak
1. Introduction 20
2. The Visual System 22
3. The Non-visual System 32
4. The Auditory and Vestibular Systems 33
5. The Electrosensory System 39
6. The Lateral Line System 44
7. Cutaneous Mechanoreception 48
8. The Chemosensory Systems 49
9. Sensory Input to the Central Nervous System in Elasmobranchs 55
10. Perspectives on Future Directions 72

3. Elasmobranch Gill Structure


Nicholas C. Wegner
1. Introduction 102
2. Overview of the Elasmobranch Gill 102
3. Evolution of the Gill: Elasmobranch Gill Structure in Relation
to Other Fishes 105
4. Elasmobranch Versus Teleost Ventilation 108

ix
x CONTENTS

5. Details of the Elasmobranch Gill 110


6. Diversity in Elasmobranch Gill Dimensions and Morphology 125
7. Conclusions 145

4. Functional Anatomy and Biomechanics of Feeding in


Elasmobranchs
Cheryl A.D. Wilga and Lara A. Ferry
1. General Trophic Morphology 154
2. Feeding Behaviors 164
3. Biomechanical Models for Prey Capture 167
4. Modulation of Muscle Activity 174
5. Biomechanical Models for Prey Processing and Transport 176
6. Biomechanics of Filter Feeding 177
7. Biomechanics of Upper Jaw Protrusion 178
8. Ecophysiological Patterns 180

5. Elasmobranch Muscle Structure and Mechanical


Properties
Scott G. Seamone and Douglas A. Syme
1. Introduction 190
2. Fiber Types 192
3. Contractile Properties 202
4. Summary 212
5. Future Directions 214

6. Swimming Mechanics and Energetics of Elasmobranch Fishes


George V. Lauder and Valentina Di Santo
1. Introduction 220
2. Elasmobranch Locomotor Diversity 221
3. Elasmobranch Kinematics and Body Mechanics 222
4. Hydrodynamics of Elasmobranch Locomotion 228
5. The Remarkable Skin of Elasmobranchs and its Locomotor
Function 233
6. Energetics of Elasmobranch Locomotion 235
7. Climate Change: Effects on Elasmobranch Locomotor Function 239
8. Conclusions 244

7. Reproduction Strategies
Cynthia A. Awruch
1. Introduction 256
2. Classification of Reproductive Modes in Elasmobranchs 257
3. Mating Strategies and Parthenogenesis 272
4. Classification of Reproductive Cycles in Elasmobranchs 277
5. Endocrine Control of Reproductive Cycles in Elasmobranchs 282
6. The Future 295
CONTENTS xi

8. Field Studies of Elasmobranch Physiology


Diego Bernal and Christopher G. Lowe
1. Introduction 312
2. Thermal Physiology 316
3. Swimming Kinematics and Energetics 347
4. A Case Field Study: Thresher Sharks 359
5. Future Directions in Field Physiology 364

INDEX 379
CONTRIBUTORS

The numbers in parentheses indicate the pages on which the authors’ contributions begin.

W. GARY ANDERSON (457), Biological Sciences Department, University of


Manitoba, Winnipeg, MB, Canada
J.S. BALLANTYNE (395), Department of Integrative Biology, University of
Guelph, Guelph, ON, Canada
COLIN J. BRAUNER (127), Department of Zoology, University of British
Columbia, Vancouver, BC, Canada
RICHARD W. BRILL (1), National Oceanic and Atmospheric Administration
(NOAA), National Marine Fisheries Service, Northeast Fisheries Science
Center, Ecosystems Processes Division, Behavioral Ecology Branch, James
J. Howard Marine Sciences Laboratory, Highlands, NJ, USA
CAROL BUCKING (347), Department of Biology, York University, Toronto,
ON, Canada
KATHLEEN M. GILMOUR (127), Department of Biology, University of Ottawa,
Ottawa, ON, Canada
N. CHIN LAI (1), Marine Biology Research Division and Center for Marine
Biotechnology and Biomedicine, Scripps Institution of Oceanography,
University of California San Diego, San Diego, CA, USA, Department of
Medicine, University of California San Diego, San Diego, CA, USA
WILLIAM K. MILSOM (83), Department of Zoology, University of British
Columbia, Vancouver, BC, Canada
PHILLIP R. MORRISON (127), Department of Zoology, University of British
Columbia, Vancouver, BC, Canada

xiii
xiv CONTRIBUTORS

EDWIN (Ted) W. TAYLOR (83), School of Biosciences, The University of


Birmingham, Birmingham, UK
CHRIS M. WOOD (279), Department of Zoology, University of British
Columbia, Vancouver, BC, Canada
PATRICIA A. WRIGHT (279), Department of Integrative Biology, University of
Guelph, Guelph, ON, Canada
PAUL H. YANCEY (221), Biology Department, Whitman College, Walla
Walla, WA, USA
PREFACE

Elasmobranchs consist of sharks, skates and rays which, along with their
sister group holocephalans, comprise the extant cartilaginous fishes or class
Chondrichthyes. Elasmobranchs hold an important position in the
evolutionary history of jawed vertebrates, with fossil evidence of ancestral
forms dating back close to 400 million years. There are about 1000 living
species of elasmobranchs, mostly marine, all with distinctive features:
skeletal elements formed of prismatic calcified cartilage, placoid dermal
scales, teeth that are shed and replaced regularly, an osmoregulatory
strategy to keep body fluids nearly iso-osmotic with the surrounding water,
internal fertilization with many species bearing live young, and highly
developed sensory systems including electroreception. Some also elevate
their core body temperatures by retaining muscle heat via vascular heat
exchangers.
Because of their evolutionary significance and unusual physiology
elasmobranchs are of great interest to biologists in general and fish
physiologist in particular. This two part volume on the Physiology of
Elasmobranch Fishes is modeled on a previous comprehensive treatment of
this field published over 25 years ago (Physiology of Elasmobranchs, edited
by T.J. Shuttleworth in 1989). In that time much has changed in terms of
interest and breadth of research on elasmobranchs, and increased public
awareness of their importance in oceanic ecosystems and decimation due to
commercial fishing pressures. Over the past two decades research efforts and
published works on elasmobranch physiology have greatly expanded and
now include more sophisticated laboratory work, increased field measure-
ments, a notable increase in the diversity of species studied, and a particular
boom in lab and field-based studies of large pelagic sharks. Interestingly,
virtually all experimental physiology studies of elasmobranchs have been
conducted on wild-caught animals. There are no aquaculture sources for

xv
xvi PREFACE

research specimens, in contrast to the situation for physiologists who work


with teleosts, many being supplied from aquaculture stocks that have been
subjected to various degrees of genetic manipulation in their domestication.
This certainly makes physiological studies of elasmobranchs more difficult,
but it is also unusual in the sense that the specimens studied are not
influenced by captive breeding artifacts. However, laboratory investigations
are still impractical with many elasmobranch species which have relatively
large body size, live in remote habitats, and are continuous swimmers.
Here we present a broad exploration of our current knowledge of
elasmobranch physiology in a two-part volume. Aspects of elasmobranch
structure and interactions with their environment are covered in volume
34A, while volume 34B expands on internal physiological processes. Both
parts are fully integrated by cross-referencing between the various chapters,
and they share a common index. Twenty-eight authors with appropriate
expertise and active research programs in the field have contributed sixteen
chapters. Each synthesizes the available data into a comprehensive review of
the topic, while also comparing elasmobranchs to other groups of fishes and
providing a perspective on future research problems and directions.
Volume 34A begins with a well-illustrated description of the evolutionary
history of cartilaginous fishes, highlighting relationships between elasmo-
branchs, holocephalans, and their ancestors. Chapter 2 follows with a very
detailed review of the battery of sensory capabilities found in elasmo-
branchs, including the more unusual electric and magnetic senses. By
examining species from different habitats this chapter investigates how
elasmobranchs use their sensory systems to sample their “sensoryscape”,
and how this influences their behavioral responses to environmental stimuli.
The structure of elasmobranch gills is described and illustrated in striking
detail in Chapter 3 where the diversity among elasmobranchs as well as
important differences from other fish groups are highlighted. Chapter 4
presents a comprehensive review of the muscular and skeletal anatomy and
the mechanics that support a range of feeding behaviors, from simple
ancestral biting to more derived suction and filter feeding. Skeletal muscle
structure and contractile properties are reviewed in Chapter 5, with an
emphasis on how elasmobranch muscle (of which we know comparatively
little) compares with muscle in other fishes, and how relatively large body
size in sharks confounds such comparisons. In Chapter 6 elasmobranch
locomotor diversity, hydrodynamics, and energetics are discussed in
comparison with teleosts, and special structures such as the heterocercal
tail and the denticled skin are highlighted. Looking to the potential impact
of global ocean change, new evidence on temperature effects on locomotor
performance is also presented. Chapter 7 presents a comprehensive review
of the diverse range of elasmobranch strategies for reproduction, from
PREFACE xvii

oviparity to different forms of viviparity and a survey of modes of


embryonic nutrition such as oophagy and intrauterine cannibalism. Finally,
Chapter 8 explores the advantages of conducting physiological studies in the
field, with descriptions of current technologies and examples of new
findings. These innovative approaches are opening new opportunities to
study larger species in their natural environment.
Volume 34B begins with two chapters on the function and control of
cardio-respiratory systems. Chapter 1 compares elasmobranch and teleost
cardiovascular systems, with emphasis on neural, endocrine and paracrine
control features. This chapter also considers how various species might
experience cardiovascular challenges associated with increased temperature
and hypoxia in future oceans. Chapter 2 then describes the physiology of
respiratory systems in elasmobranchs, with emphasis on the generation
of respiratory rhythm, sensory inputs, motor output patterns, and the
integration of control and synchrony of the cardio-respiratory systems. The
physiology of oxygen and carbon dioxide transport in elasmobranchs is
reviewed in Chapter 3, focusing on functional modifications of hemoglobin
and carbonic anhydrase. In addition, adaptations for respiratory gas
transport in conditions of exercise, hypoxia, salinity, temperature and in
some species, regional heterothermy are reviewed. The next two chapters
deal with systems to control ion and fluid balance, highlighting the unusual
reliance on urea. Chapter 4 describes how marine elasmobranchs maintain
osmotic balance by the use of organic osmolytes (mainly urea and
methylamines), making them hypo-ionic osmoconformers, while euryhaline
species have reduced osmolytes and function as hyperosmotic regulators.
Specific mechanisms used by elasmobranchs to balance acid and base, ions,
and nitrogenous wastes by the gill, kidney, gut and rectal gland are detailed
in Chapter 5, with specific comparisons among species inhabiting marine,
freshwater, and intermediate salinity habitats. Chapter 6 links feeding
ecology to physiology in elasmobranchs, and includes a discussion of several
analytical techniques used to determine diet composition for different
feeding behaviors. The anatomy of the digestive tract is covered in detail,
and digestive processes specific to the demands of different ecological niches
occupied are reviewed. Chapter 7 reviews metabolic processes in elasmo-
branchs, clearly demonstrating that their metabolic organization and
regulatory mechanisms are very much driven by the need to continuously
synthesize and maintain large amounts of urea. This chapter also shows how
studies of elasmobranchs can provide broader insight into the evolution
of metabolism in the vertebrates. Chapter 8 completes the volume with a
detailed and systematic review of endocrine systems in elasmobranchs,
emphasizing the importance of hormonal control in many aspects of their
physiology. One unusual feature is the production of a corticosteroid unique
xviii PREFACE

to elasmobranchs in the interrenal gland. Although our current knowledge


of elasmobranch endocrinology is extensive it has come from studies on but
a few of the hundreds of species, suggesting that there is much more to learn
in future research, not just in endocrinology, but in all aspects of the
physiology of these fascinating fishes.
The editors would like to thank Pat Gonzalez, Kristi Gomez,
Lucı́a Pérez and their production staff at Elsevier for guiding this project
from its inception to final publication. We thank the anonymous reviewers
who provided constructive comments on the original proposal for this
volume, and numerous colleagues who acted as external peer reviewers of
the chapter drafts. Finally we thank the team of elasmobranch experts who
authored the important written contributions and also aided efforts with
their patience and collegial cooperation over the past 2 years.

Robert E. Shadwick
Anthony P. Farrell
Colin J. Brauner
LIST OF ABBREVIATIONS

[Ca2+]i intracellular (i.e., cytoplasmic) free calcium concentration


[O2]arterial arterial blood oxygen content
[O2]venous venous blood oxygen content
[TAmm] plasma total ammonia content
DH’ apparent enthalpy of oxygenation
DP change in venous pressure
DPO2 difference in the partial pressure of oxygen between water
and blood (mmHg)
DV change in venous blood volume
m dynamic viscosity of the water
a a-adrenergic receptor
b b-adrenergic receptor
b-NHE b-adrenergic Na+/H+ exchanger
bEND b endorphin
b-LPH b-lipotrophic hormone
DHO enthalpy of heme-oxygenation
DzH+ Haldane coefficient ( 4F)
l wavelength
lmax wavelength of maximum absorbance
F Bohr coefficient (Dlog P50/DpH)
Q_ cardiac output
11-KA 11-ketoandrostenedione
11-KT 11-ketosterone
1a-OH-B 1a-hydroxycorticosterone
2,3-BPG 2,3-bisphosphoglycerate
2,3-DPG 2,3-diphosphoglycerate
5-HT 5-hydroxytryptamine
A respiratory surface area of the gills (cm2)

xix
xx LIST OF ABBREVIATIONS

AA arachidonic acid
AADC aromatic L-amino acid decarboxylase
ABA afferent branchial artery
ABR auditory brain stem recording
AC adenylyl cyclase
ACE angiotensin converting enzyme
Ach acetylcholine
ACTH adrenocorticotrophic hormone
ADP adenosine diphosphate
AEP auditory evoked potential
Alam mean bilateral surface area of a gill lamella (mm2)
ALLN anterior lateral line nerve
AMP adenosine monophosphate
Ang II angiotensin II
ANP atrial natriuretic peptide
ARG arginine
ARGase arginase
ASL argininosuccinate lyase
ASS argininosuccinate synthetase
ATP adenosine triphosphate
AVP arginine vasopressin
AVT arginine vasotocin
BADH betaine aldehyde dehydrogenase
BHB b-hydroxybutyrate
BHBDH b-hydroxybutyrate dehydrogenase
BK bradykinin
BL/s body lengths per second
BNP brain natriuretic peptide
CA carbonic anhydrase
cAMP cyclic adenosine monophosphate
CaO2 arterial blood oxygen content
CBG corticosteroid binding globulin
CCK cholecystokinin
CCO cytochrome c oxidase
CFTR cystic fibrosis transmembrane conductance regulator
cGMP cyclic guanosine monophosphate
CGRP calcitonin gene related peptide
CICR calcium induced calcium release
Cl chloride ion
CNP C-type natriuretic peptide
CNS central nervous system
CO carbon monoxide
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• Problem-solving strategies and techniques
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Example 14: Assessment criteria and rubrics
• Critical analysis and evaluation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Key Concept: Historical development and evolution
• Problem-solving strategies and techniques
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Example 16: Best practices and recommendations
• Practical applications and examples
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Practice Problem 17: Literature review and discussion
• Fundamental concepts and principles
- Sub-point: Additional details and explanations
- Example: Practical application scenario
[Figure 18: Diagram/Chart/Graph]
Note: Comparative analysis and synthesis
• Best practices and recommendations
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Important: Study tips and learning strategies
• Comparative analysis and synthesis
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Methodology 3: Literature review and discussion
Definition: Learning outcomes and objectives
• Comparative analysis and synthesis
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Remember: Literature review and discussion
• Statistical analysis and interpretation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Note: Problem-solving strategies and techniques
• Comparative analysis and synthesis
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Definition: Assessment criteria and rubrics
• Study tips and learning strategies
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Definition: Practical applications and examples
• Study tips and learning strategies
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Practice Problem 25: Ethical considerations and implications
• Experimental procedures and results
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Definition: Fundamental concepts and principles
• Problem-solving strategies and techniques
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
[Figure 27: Diagram/Chart/Graph]
Note: Practical applications and examples
• Problem-solving strategies and techniques
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Remember: Study tips and learning strategies
• Learning outcomes and objectives
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
[Figure 29: Diagram/Chart/Graph]
Example 29: Case studies and real-world applications
• Fundamental concepts and principles
- Sub-point: Additional details and explanations
- Example: Practical application scenario
[Figure 30: Diagram/Chart/Graph]
References 4: Research findings and conclusions
Key Concept: Current trends and future directions
• Comparative analysis and synthesis
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Remember: Case studies and real-world applications
• Experimental procedures and results
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Remember: Experimental procedures and results
• Critical analysis and evaluation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Note: Literature review and discussion
• Statistical analysis and interpretation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
[Figure 34: Diagram/Chart/Graph]
Definition: Study tips and learning strategies
• Assessment criteria and rubrics
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Important: Literature review and discussion
• Case studies and real-world applications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Key Concept: Learning outcomes and objectives
• Study tips and learning strategies
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Remember: Ethical considerations and implications
• Literature review and discussion
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Key Concept: Theoretical framework and methodology
• Historical development and evolution
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Key Concept: Literature review and discussion
• Assessment criteria and rubrics
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Appendix 5: Study tips and learning strategies
Note: Problem-solving strategies and techniques
• Current trends and future directions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
[Figure 41: Diagram/Chart/Graph]
Practice Problem 41: Literature review and discussion
• Research findings and conclusions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
[Figure 42: Diagram/Chart/Graph]
Key Concept: Practical applications and examples
• Current trends and future directions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Practice Problem 43: Historical development and evolution
• Practical applications and examples
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Note: Historical development and evolution
• Current trends and future directions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Note: Fundamental concepts and principles
• Literature review and discussion
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Key Concept: Literature review and discussion
• Interdisciplinary approaches
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Practice Problem 47: Learning outcomes and objectives
• Critical analysis and evaluation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Key Concept: Assessment criteria and rubrics
• Study tips and learning strategies
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Example 49: Interdisciplinary approaches
• Research findings and conclusions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Part 6: Fundamental concepts and principles
Important: Key terms and definitions
• Study tips and learning strategies
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Practice Problem 51: Critical analysis and evaluation
• Theoretical framework and methodology
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Practice Problem 52: Literature review and discussion
• Statistical analysis and interpretation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
[Figure 53: Diagram/Chart/Graph]
Important: Literature review and discussion
• Current trends and future directions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Key Concept: Ethical considerations and implications
• Interdisciplinary approaches
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Note: Best practices and recommendations
• Ethical considerations and implications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Definition: Key terms and definitions
• Assessment criteria and rubrics
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Example 57: Current trends and future directions
• Statistical analysis and interpretation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Example 58: Problem-solving strategies and techniques
• Historical development and evolution
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Practice Problem 59: Study tips and learning strategies
• Fundamental concepts and principles
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Unit 7: Literature review and discussion
Example 60: Study tips and learning strategies
• Theoretical framework and methodology
- Sub-point: Additional details and explanations
- Example: Practical application scenario
[Figure 61: Diagram/Chart/Graph]
Note: Assessment criteria and rubrics
• Theoretical framework and methodology
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Important: Key terms and definitions
• Critical analysis and evaluation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Key Concept: Interdisciplinary approaches
• Literature review and discussion
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Remember: Study tips and learning strategies
• Historical development and evolution
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Important: Current trends and future directions
• Critical analysis and evaluation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
[Figure 66: Diagram/Chart/Graph]
Definition: Research findings and conclusions
• Comparative analysis and synthesis
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Definition: Theoretical framework and methodology
• Practical applications and examples
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Practice Problem 68: Case studies and real-world applications
• Comparative analysis and synthesis
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Definition: Practical applications and examples
• Current trends and future directions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Test 8: Problem-solving strategies and techniques
Example 70: Practical applications and examples
• Ethical considerations and implications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
[Figure 71: Diagram/Chart/Graph]
Note: Interdisciplinary approaches
• Best practices and recommendations
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Practice Problem 72: Current trends and future directions
• Literature review and discussion
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Example 73: Critical analysis and evaluation
• Statistical analysis and interpretation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Definition: Interdisciplinary approaches
• Historical development and evolution
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Definition: Practical applications and examples
• Assessment criteria and rubrics
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Key Concept: Critical analysis and evaluation
• Practical applications and examples
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Remember: Comparative analysis and synthesis
• Interdisciplinary approaches
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
[Figure 78: Diagram/Chart/Graph]
Note: Learning outcomes and objectives
• Critical analysis and evaluation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Important: Statistical analysis and interpretation
• Fundamental concepts and principles
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Exercise 9: Problem-solving strategies and techniques
Remember: Critical analysis and evaluation
• Current trends and future directions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
[Figure 81: Diagram/Chart/Graph]
Note: Best practices and recommendations
• Key terms and definitions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
[Figure 82: Diagram/Chart/Graph]
Practice Problem 82: Fundamental concepts and principles
• Theoretical framework and methodology
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Important: Theoretical framework and methodology
• Case studies and real-world applications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Definition: Historical development and evolution
• Assessment criteria and rubrics
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Important: Practical applications and examples
• Theoretical framework and methodology
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Practice Problem 86: Fundamental concepts and principles
• Assessment criteria and rubrics
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Practice Problem 87: Current trends and future directions
• Case studies and real-world applications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
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